Demystifying ecological connectivity for actionable spatial conservation planning
Authors: Maria Beger, Anna Metaxas, Arieanna C. Balbar, Jennifer A. McGowan, Remi Daigle, Caitlin D. Kuempel, Eric A. Treml, Hugh P. Possingham
Journal: Trends in Ecology & Evolution (2022)

There is a disconnect between global high-level conservation goals and on-the-ground actions such as maintaining ecosystem services or persistence and local planning of protected areas.Dynamic processes such as ecological connectivity underpin species persistence and ecosystem resilience but are difficult to represent in mathematical spatial planning problems for protected areas.Quantitative and SMART (specific – measurable – action-oriented – realistic – time-bound) conservation objectives can provide a link between high-level conservation goals and local or regional design and implementation of functionally connected protected area networks.With current implementation gaps of protected area commitments and increasing climate change threats, there is tremendous opportunity to use quantifiable objectives for ecological connectivity as a vehicle to future-proof protected area networks to help achieve global conservation goals. Connectivity underpins the persistence of life; it needs to inform biodiversity conservation decisions. Yet, when prioritising conservation areas and developing actions, connectivity is not being operationalised in spatial planning. The challenge is the translation of flows associated with connectivity into conservation objectives that lead to actions. Connectivity is nebulous, it can be abstract and mean different things to different people, making it difficult to include in conservation problems. Here, we show how connectivity can be included in mathematically defining conservation planning objectives. We provide a path forward for linking connectivity to high-level conservation goals, such as increasing species’ persistence. We propose ways to design spatial management areas that gain biodiversity benefit from connectivity. Connectivity underpins the persistence of life; it needs to inform biodiversity conservation decisions. Yet, when prioritising conservation areas and developing actions, connectivity is not being operationalised in spatial planning. The challenge is the translation of flows associated with connectivity into conservation objectives that lead to actions. Connectivity is nebulous, it can be abstract and mean different things to different people, making it difficult to include in conservation problems. Here, we show how connectivity can be included in mathematically defining conservation planning objectives. We provide a path forward for linking connectivity to high-level conservation goals, such as increasing species’ persistence. We propose ways to design spatial management areas that gain biodiversity benefit from connectivity. In a world of dwindling natural resources and increasing human pressures, global conservation goals aim to ensure that habitats and species can persist into the future. Most notably, the United Nations Sustainable Development Goals (SDG) SDG14 (life below the water) and SDG15 (life on land), and the Convention on Biological Diversity’s (CBD) post-2020 Global Biodiversity Framework aim to halt loss of biodiversity and associated ecosystem services. A dominant mechanism to achieve these goals will be through area-based conservation and management [1.Pressey R.L. et al.The mismeasure of conservation.Trends Ecol. Evol. 2021; 36: 808-821Abstract Full Text Full Text PDF PubMed Scopus (28) Google Scholar, 2.Garibaldi L.A. et al.Working landscapes need at least 20% native habitat.Conserv. Lett. 2021; 14e12773Crossref Scopus (109) Google Scholar, 3.Nicholson E. et al.Scientific foundations for an ecosystem goal, milestones and indicators for the post-2020 global biodiversity framework.Nat. Ecol. Evol. 2021; 5: 1338-1349Crossref PubMed Scopus (42) Google Scholar], with specific goals of achieving the protection of ‘well-connected systems’. Connectivity (see Glossary) underpins the persistence of populations, species, communities, and ecosystems, and thus needs to play a pivotal role in conservation strategies (e.g., [4.Wood S.L.R. et al.Missing interactions: the current state of multispecies connectivity analysis.Front. Ecol. Evol. 2022; 10830822Crossref Scopus (7) Google Scholar, 5.Magris R.A. et al.Biologically representative and well-connected marine reserves enhance biodiversity persistence in conservation planning.Conserv. Lett. 2018; 11e12439Crossref Scopus (71) Google Scholar, 6.Riginos C. Beger M. Incorporating genetic measures of connectivity and adaptation in marine spatial planning for corals.in: van Oppen M. Aranda Lastra M.I. Coral Reef Conservation and Restoration in the ‘Omics’ Age. Springer, 2022Crossref Google Scholar]). Yet, conceptual advancements and tools to quantitatively integrate connectivity for and across land, freshwater, and marine systems with area-based conservation are still being developed (e.g., [5.Magris R.A. et al.Biologically representative and well-connected marine reserves enhance biodiversity persistence in conservation planning.Conserv. Lett. 2018; 11e12439Crossref Scopus (71) Google Scholar,7.Tulloch V.J.D. et al.Minimizing cross-realm threats from land-use change: a national-scale conservation framework connecting land, freshwater and marine systems.Biol. Conserv. 2021; 254108954Crossref Scopus (13) Google Scholar, 8.Hermoso V. et al.Conservation planning across realms: enhancing connectivity for multi-realm species.J. Appl. Ecol. 2021; 58: 644-654Crossref Scopus (10) Google Scholar, 9.Daigle R. et al.Operationalizing ecological connectivity in spatial conservation planning with Marxan Connect.Methods Ecol. Evol. 2020; 11: 570-579Crossref Scopus (52) Google Scholar, 10.Heino J. et al.Integrating dispersal proxies in ecological and environmental research in the freshwater realm.Environ. Rev. 2017; 25: 334-349Crossref Scopus (81) Google Scholar]), and are only implemented in a fraction of existing conservation areas [11.Balbar A.C. Metaxas A. The current application of ecological connectivity in the design of marine protected areas.Global Ecol. Conserv. 2019; 17e00569PubMed Google Scholar,12.Ward M. et al.Just ten percent of the global terrestrial protected area network is structurally connected via intact land.Nat. Commun. 2020; 11: 4563Crossref PubMed Scopus (87) Google Scholar]. In this opinion article, we define connectivity as the flow of energy, materials, and organisms across space. At the species level, this connectivity includes adult and propagule dispersal, species movement and migration, species interactions, and ontogenetic linkages. Flow processes of energy, materials, and organisms that underpin connectivity are dynamic, variable, and often spatially unconstrained (Box 1), generating a considerable challenge for formulating both suitable metrics and useful objectives for traditional conservation planning approaches [9.Daigle R. et al.Operationalizing ecological connectivity in spatial conservation planning with Marxan Connect.Methods Ecol. Evol. 2020; 11: 570-579Crossref Scopus (52) Google Scholar,13.Keeley A.T.H. et al.Connectivity metrics for conservation planning and monitoring.Biol. Conserv. 2021; 255109008Crossref Scopus (41) Google Scholar,14.Jafari N. et al.Achieving full connectivity of sites in the multiperiod reserve network design problem.Comput. Oper. Res. 2017; 81: 119-127Crossref Scopus (16) Google Scholar]. The variable characteristics and scale of flow processes have led to diverse characterisations of connectivity in environmental conservation, ranging from spatial wetland linkages for amphibians [15.Heard G.W. et al.Refugia and connectivity sustain amphibian metapopulations afflicted by disease.Ecol. Lett. 2015; 18: 853-863Crossref PubMed Scopus (65) Google Scholar] to recent genetic exchange among populations [16.Xuereb A. et al.Individual-based eco-evolutionary models for understanding adaptation in changing seas.Proc. R. Soc. Lond. Ser. B Biol. Sci. 2021; 288: 20212006PubMed Google Scholar] (Table 1). Assessments of the global protected area estate highlight shortfalls in capturing dynamic ecological processes, such as connectivity, where only 9.7% of intact land is protected and connected [12.Ward M. et al.Just ten percent of the global terrestrial protected area network is structurally connected via intact land.Nat. Commun. 2020; 11: 4563Crossref PubMed Scopus (87) Google Scholar], two thirds of critical areas for the flow of animals on land are not conserved [17.Brennan A. et al.Functional connectivity of the world’s protected areas.Science. 2022; 376: 1101-1104Crossref PubMed Scopus (35) Google Scholar], only 17% of the world’s free-flowing rivers are protected [18.Opperman J.J. et al.Safeguarding free-flowing rivers: the global extent of free-flowing rivers in protected areas.Sustainability. 2021; 13: 2805Crossref Scopus (7) Google Scholar], and 90.5% of marine species have less than 5% of their ranges protected [19.Klein C.J. et al.Shortfalls in the global protected area network at representing marine biodiversity.Sci. Rep. 2015; 5: 17539Crossref PubMed Scopus (112) Google Scholar]. This implementation gap is often because broad conservation goals for connectivity are difficult to translate into quantitative conservation objectives, data that measure connectivity are difficult to acquire, and there is no scientific consensus on the appropriate metrics to use to assess connectivity retention or improvement [13.Keeley A.T.H. et al.Connectivity metrics for conservation planning and monitoring.Biol. Conserv. 2021; 255109008Crossref Scopus (41) Google Scholar], especially for multiple species [4.Wood S.L.R. et al.Missing interactions: the current state of multispecies connectivity analysis.Front. Ecol. Evol. 2022; 10830822Crossref Scopus (7) Google Scholar].Box 1Types and scales of connectivity that hinder its estimationA key hurdle to including connectivity in spatial planning is its spatial–temporal complexity. The directionality, spatial constraint, and spatial–temporal scales of the flows of energy, materials, and organisms vary with their physical or ecological process, the properties of the environment, and the flowing entity (Figure I). These flows can occur in any medium (e.g., land, river, ocean, air) and across spatial scales ranging from metres to across continents, hemispheres, or ocean basins. Ensuing connectivity may be manifested and relevant over time scales ranging from hours to centuries or even longer (as in the case of evolutionary time scales). Many flows can be either symmetrical (e.g., movement along animal migration corridors) or asymmetrical (e.g., movement across ontogeny, seed or larva dispersal). This variability underpins the measurements of connectivity that are appropriate in each case.Directed flows involve the movement of an entity along a single, dominant direction (Figure II). These flows can be constrained, with relatively low lateral variation in the path (e.g., upstream or downstream salmon migration, downstream transport of leaf litter,movement along terrestrial migration corridors, annual bird migrations across continents or ocean basins). Directed flows can be unconstrained when lateral variation in the path is high. This variation can result from the movement of the medium or of the moving entity, for example in spread of invasive/range-expanding species along a coast with a boundary current, turtle migration from foraging to spawning grounds, or ungulate migration across seasonal feeding grounds.In diffuse flows, movement proceeds along a number of directions, and can originate from a single source (e.g., during an oil spill, foraging from a nesting aggregation) or multiple sources (e.g., multiple introductions of non-native species) (Figure III). They can also be either constrained with relatively clear movement corridors or pathways (e.g., detrital dispersal into valleys or basins, foraging within a particular ambit, spread of invasive species or disease within a bounded suitable habitat) or unconstrained with multiple possible pathways such as the movement of propagules that are dispersed by wind or ocean current.Figure IIDirected connectivity has a dominant direction of movement and is easier to conceptualise.View Large Image Figure ViewerDownload Hi-res image Download (PPT)Figure IIIDiffuse connectivity is mixed in direction and strength and is extremely difficult to estimate.View Large Image Figure ViewerDownload Hi-res image Download (PPT)Table 1Connectivity as a value-laden concept. Selected contexts of connectivity and potential audiences applying these concepts for spatial conservation area network planningType of connectivityDefinition/examplesReference for definitionExample user groupLand–sea connectivityFlows of sediment and pollutants from rivers into the sea, and movement of animals between land, rivers, and the sea[70.Suárez-Castro A.F. et al.Global forest restoration opportunities to foster coral reef conservation.Glob. Chang. Biol. 2021; 27: 5238-5252Crossref PubMed Scopus (12) Google Scholar]Ecologist, environmental scientist, engineerOntogenetic connectivityMovement of individuals occurring as part of life cycles (metres to thousands of km), e.g., amphibians[15.Heard G.W. et al.Refugia and connectivity sustain amphibian metapopulations afflicted by disease.Ecol. Lett. 2015; 18: 853-863Crossref PubMed Scopus (65) Google Scholar,48.Kot C.Y. et al.Network analysis of sea turtle movements and connectivity: a tool for conservation prioritization.Divers. Distrib. 2022; 28: 810-829Crossref Scopus (8) Google Scholar]Ecologist, park managerCorridorsDistinct habitant patches are linked such that movement of animals can be facilitated. Disruption of corridors often occurs due to fragmentation[36.Keeley A.T.H. et al.Thirty years of connectivity conservation planning: an assessment of factors influencing plan implementation.Environ. Res. Lett. 2019; 14103001Crossref Scopus (55) Google Scholar]Environmental scientist, wildlife biologist, park manager, tourism operatorPathogen dispersalAirborne dispersal of fungal spores (regional and continental scale, 50–5000 km)[46.Meyer M. et al.Quantifying airborne dispersal routes of pathogens over continents to safeguard global wheat supply.Nat. Plants. 2017; 3: 780-786Crossref PubMed Scopus (55) Google Scholar]EpidemiologistPollutant advection and diffusionTransport of pollutants in a medium (e.g., oil spill, sewage transport in water)[54.Chaturvedi S.K. et al.An assessment of oil spill detection using Sentinel 1 SAR-C images.J. Ocean Eng. Sci. 2020; 5: 116-135Crossref Scopus (40) Google Scholar]Engineer, geophysicistDispersal connectivityThe movement of propagules or juveniles among spatially distinct habitat patches. Scale highly variable, dependent on medium and species[55.Hüssy K. et al.Trace element patterns in otoliths: the role of biomineralization.Rev. Fish. Sci. Aquacult. 2021; 29: 445-477Crossref Scopus (79) Google Scholar,57.Lett C. et al.Converging approaches for modeling the dispersal of propagules in air and sea.Ecol. Model. 2020; 415108858Crossref Scopus (5) Google Scholar,58.Cecino G. Treml E.A. Local connections and the larval competency strongly influence marine metapopulation persistence.Ecol. Appl. 2021; 31e02302Crossref PubMed Scopus (6) Google Scholar,79.Harrison H.B. et al.A connectivity portfolio effect stabilizes marine reserve performance.Proc. Natl. Acad. Sci. U. S. A. 2020; 117: 25595Crossref PubMed Scopus (38) Google Scholar]Modeller, hydrodynamics engineer, oceanographer, ecologistMigrationThe scheduled movement of individuals[47.Schuster R. et al.Optimizing the conservation of migratory species over their full annual cycle.Nat. Commun. 2019; 10: 1754Crossref PubMed Scopus (47) Google Scholar,83.Somveille M. et al.A general theory of avian migratory connectivity.Ecol. Lett. 2021; 24: 1848-1858Crossref PubMed Scopus (13) Google Scholar]Wildlife biologist, ornithologist, park manager, tourism operatorGenetic connectivityThe movement of genetic material between nearby or distant habitat regions over multiple generations[16.Xuereb A. et al.Individual-based eco-evolutionary models for understanding adaptation in changing seas.Proc. R. Soc. Lond. Ser. B Biol. Sci. 2021; 288: 20212006PubMed Google Scholar]Geneticist, evolutionary ecologistTemporal connectivityLinkages among sites as species shift their ranges over time[51.Williams S.H. et al.Incorporating connectivity into conservation planning for the optimal representation of multiple species and ecosystem services.Conserv. Biol. 2020; 34: 934-942Crossref PubMed Scopus (13) Google Scholar,84.Makino A. et al.Spatio-temporal marine conservation planning to support high-latitude coral range expansion under climate change.Divers. Distrib. 2014; 2014: 6-12Google Scholar]Climate scientist, global change ecologistEnergy flowTransport of nutrients as part of animal movement[39.Venarsky M.P. et al.Spatial and temporal variation of fish community biomass and energy flow throughout a tropical river network.Freshw. Biol. 2020; 65: 1782-1792Crossref Scopus (12) Google Scholar]Ecologist, chemist Open table in a new tab A key hurdle to including connectivity in spatial planning is its spatial–temporal complexity. The directionality, spatial constraint, and spatial–temporal scales of the flows of energy, materials, and organisms vary with their physical or ecological process, the properties of the environment, and the flowing entity (Figure I). These flows can occur in any medium (e.g., land, river, ocean, air) and across spatial scales ranging from metres to across continents, hemispheres, or ocean basins. Ensuing connectivity may be manifested and relevant over time scales ranging from hours to centuries or even longer (as in the case of evolutionary time scales). Many flows can be either symmetrical (e.g., movement along animal migration corridors) or asymmetrical (e.g., movement across ontogeny, seed or larva dispersal). This variability underpins the measurements of connectivity that are appropriate in each case. Directed flows involve the movement of an entity along a single, dominant direction (Figure II). These flows can be constrained, with relatively low lateral variation in the path (e.g., upstream or downstream salmon migration, downstream transport of leaf litter,movement along terrestrial migration corridors, annual bird migrations across continents or ocean basins). Directed flows can be unconstrained when lateral variation in the path is high. This variation can result from the movement of the medium or of the moving entity, for example in spread of invasive/range-expanding species along a coast with a boundary current, turtle migration from foraging to spawning grounds, or ungulate migration across seasonal feeding grounds. In diffuse flows, movement proceeds along a number of directions, and can originate from a single source (e.g., during an oil spill, foraging from a nesting aggregation) or multiple sources (e.g., multiple introductions of non-native species) (Figure III). They can also be either constrained with relatively clear movement corridors or pathways (e.g., detrital dispersal into valleys or basins, foraging within a particular ambit, spread of invasive species or disease within a bounded suitable habitat) or unconstrained with multiple possible pathways such as the movement of propagules that are dispersed by wind or ocean current.Figure IIIDiffuse connectivity is mixed in direction and strength and is extremely difficult to estimate.View Large Image Figure ViewerDownload Hi-res image Download (PPT) The shortfalls in implementing connectivity in spatial management can be explained in part by the fact that the concept of connectivity is broad, complex, and means different things to different people at different scales and times. There are many different conceptualisations of ecological connectivity within the conservation community (Table 1). For example, a park manager in Kenya may be most concerned with connectivity that enhances the movement of high-value, charismatic species that bring critical tourism revenues from wildlife experiences. By contrast, a coral reef ecologist assisting with the design of marine protected areas in the Indo-Pacific may value larval connectivity, and focus conservation on reefs linked by propagule dispersal and fish spawning aggregations [20.Beger M. et al.Integrating regional conservation priorities for multiple objectives into national policy.Nat. Commun. 2015; 6: 8208Crossref PubMed Scopus (99) Google Scholar], or climate-resilient areas [21.Beyer H. et al.Risk-sensitive planning for conserving coral reefs under rapid climate change.Conserv. Lett. 2018; 11e12587Crossref Scopus (122) Google Scholar]. As is often the case in applied conservation, accounting for value-laden perspectives of different stakeholders will have trade-offs, and may hinder a unified approach to operationalise connectivity in the context of global conservation goals. One of the most widely recognised, prioritised, and historically implemented forms of connectivity on land is wildlife corridors, which connect fragmented habitats across landscapes that have been impacted by conversion or land-use change [13.Keeley A.T.H. et al.Connectivity metrics for conservation planning and monitoring.Biol. Conserv. 2021; 255109008Crossref Scopus (41) Google Scholar,22.Hilty J.A. et al.Corridor Ecology: the Science and Practice of Connectivity for Biodiversity Conservation. Island Press, 2019Google Scholar]. Habitat fragmentation affects the movement of individuals, and often, but not always [e.g., 23.Fahrig L. Ecological responses to habitat fragmentation per se.Annu. Rev. Ecol. Evol. Syst. 2017; 48: 1-23Crossref Scopus (598) Google Scholar], reduces persistence probabilities, mostly due to edge and isolation effects [24.Fletcher R.J. et al.Is habitat fragmentation good for biodiversity?.Biol. Conserv. 2018; 226: 9-15Crossref Scopus (326) Google Scholar] and by changing species interactions [25.Holyoak M. et al.Integrating disturbance, seasonality, multi-year temporal dynamics, and dormancy Into the dynamics and conservation of metacommunities.Front. Ecol. Evol. 2020; 8571130Crossref Scopus (18) Google Scholar]. However, corridor conservation, whilst important, addresses a form of structural connectivity that may serve only a few focal species, miss important and unknown barriers to movement [26.Merenlender A.M. et al.Ecological corridors for which species?.Theyra. 2022; 13: 45-55Google Scholar], and ignore essential attributes needed to retain functional connectivity, such as dynamic flows of matter and energy. By contrast, marine and freshwater conservation often focus on the functional conservation value of preserving dynamic flows in these systems [27.D'Aloia C.C. et al.Coupled networks of permanent protected areas and dynamic conservation areas for biodiversity conservation under climate change.Front. Ecol. Evol. 2019; 7: 27Crossref Scopus (48) Google Scholar, 28.Tittensor D.P. et al.Integrating climate adaptation and biodiversity conservation in the global protected ocean.Sci. Adv. 2019; 5: eaay9969Crossref PubMed Scopus (101) Google Scholar, 29.Dunn D.C. et al.The importance of migratory connectivity for global ocean policy.Proc. R. Soc. Lond. Ser. B Biol. Sci. 2019; 286: 20191472PubMed Google Scholar], but implementation in conservation plans is historically lacking [11.Balbar A.C. Metaxas A. The current application of ecological connectivity in the design of marine protected areas.Global Ecol. Conserv. 2019; 17e00569PubMed Google Scholar,18.Opperman J.J. et al.Safeguarding free-flowing rivers: the global extent of free-flowing rivers in protected areas.Sustainability. 2021; 13: 2805Crossref Scopus (7) Google Scholar]. Despite challenges, connectivity is a focal component of the CBD’s Global Biodiversity Framework and government policies for area-based conservation targets. Spatial planning as a means to achieve these targets also features prominently in ongoing discussions. Our aim is to highlight the challenges facing ‘connectivity’ as a global policy ambition and propose how high-level goals for connectivity can become quantitatively integrated into conservation plans to deliver connected conservation area networks. We recognise much progress has been made in academic research for incorporating connectivity into spatial conservation planning [8.Hermoso V. et al.Conservation planning across realms: enhancing connectivity for multi-realm species.J. Appl. Ecol. 2021; 58: 644-654Crossref Scopus (10) Google Scholar,30.Magris R.A. et al.Integrating connectivity and climate change into marine conservation planning.Biol. Conserv. 2014; 170: 207-221Crossref Scopus (138) Google Scholar, 31.Andrello M. et al.Additive effects of climate change on connectivity between marine protected areas and larval supply to fished areas.Divers. Distrib. 2015; 21: 139-150Crossref Scopus (64) Google Scholar, 32.Krueck N.C. et al.Incorporating larval dispersal into MPA design for both conservation and fisheries.Ecol. Appl. 2017; 27: 925-941Crossref PubMed Scopus (70) Google Scholar, 33.Dickson B.G. et al.Circuit-theory applications to connectivity science and conservation.Conserv. Biol. 2019; 33: 239-249Crossref PubMed Scopus (178) Google Scholar]. However, the transferability and uptake of these methods to the real-world remains limited given that these explorations often ignore objectives that are important to decision-makers on the ground (e.g., social–economic considerations, equity, political realities) [34.Virtanen E.A. et al.Marine connectivity in spatial conservation planning: analogues from the terrestrial realm.Landsc. Ecol. 2020; 35: 1021-1034Crossref Scopus (14) Google Scholar]. As a consequence, the integration of connectivity into conservation decisions by practitioners has not been fully realised even though the importance of connectivity for management goals is widely recognised, particularly for addressing threats of climate change to biodiversity and livelihoods [28.Tittensor D.P. et al.Integrating climate adaptation and biodiversity conservation in the global protected ocean.Sci. Adv. 2019; 5: eaay9969Crossref PubMed Scopus (101) Google Scholar]. Here, we provide a conceptual overview of how flows of energy, materials, and organisms, or connectivity, can support the achievement of global conservation goals. With specific examples, we illustrate how to link objectives of high-level conservation goals with local and regional connectivity objectives. Planning for area-based conservation actions (e.g., protection, restoration, or management of harvesting) that support the long-term persistence of species and ecosystem processes relates to the foundational conservation planning principle of adequacy [20.Beger M. et al.Integrating regional conservation priorities for multiple objectives into national policy.Nat. Commun. 2015; 6: 8208Crossref PubMed Scopus (99) Google Scholar,35.Kukkala A.S. Moilanen A. Core concepts of spatial prioritisation in systematic conservation planning.Biol. Rev. 2013; 88: 443-464Crossref PubMed Scopus (267) Google Scholar]. This principle ensures that the coverage and intensity of conservation actions is enough to maintain functional and adaptive structured populations or communities so they persist through time [25.Holyoak M. et al.Integrating disturbance, seasonality, multi-year temporal dynamics, and dormancy Into the dynamics and conservation of metacommunities.Front. Ecol. Evol. 2020; 8571130Crossref Scopus (18) Google Scholar,36.Keeley A.T.H. et al.Thirty years of connectivity conservation planning: an assessment of factors influencing plan implementation.Environ. Res. Lett. 2019; 14103001Crossref Scopus (55) Google Scholar,37.Jetz W. et al.Include biodiversity representation indicators in area-based conservation targets.Nat. Ecol. Evol. 2022; 6: 123-126Crossref PubMed Scopus (14) Google Scholar]. Achieving persistence requires continued functional integrity of biological communities through species interactions [38.Edelsparre A.H. et al.Habitat connectivity is determined by the scale of habitat loss and dispersal strategy.Ecol. Evol. 2018; 8: 5508-5514Crossref PubMed Scopus (15) Google Scholar] and energy flow [39.Venarsky M.P. et al.Spatial and temporal variation of fish community biomass and energy flow throughout a tropical river network.Freshw. Biol. 2020; 65: 1782-1792Crossref Scopus (12) Google Scholar,40.Benkwitt C.E. et al.Seabird nutrient subsidies alter patterns of algal abundance and fish biomass on coral reefs following a bleaching event.Glob. Chang. Biol. 2019; 25: 2619-2632Crossref PubMed Scopus (36) Google Scholar]. Flows of energy (e.g., carbon) and matter (e.g., detrital subsidies) that are critical for the persistence of ecosystems can be achieved by connectivity via animal movement and habitat linkages [41.Alberti M. et al.The complexity of urban eco-evolutionary dynamics.Bioscience. 2020; 70: 772-793Crossref Scopus (58) Google Scholar,42.Olds A.D. et al.Quantifying the conservation value of seascape connectivity: a global synthesis.Glob. Ecol. Biogeogr. 2016; 25: 3-15Crossref Scopus (107) Google Scholar]. The flow of genes amongst populations enhances their persistence by promoting genetic diversity that often underpins adaptive potential [6.Riginos C. Beger M. Incorporating genetic measures of connectivity and adaptation in marine spatial planning for corals.in: van Oppen M. Aranda Lastra M.I. Coral Ree